February 12, 2015

A story of 69 ancient Europeans

A new study on the bioRxiv includes data on 69 ancient Europeans (remember when we got excited in anticipation for the single genome of the Iceman? that was only three years ago) and adds plenty of new info to chew on for those of us interested in prehistory.

Two Near Eastern migrations into Europe
In 2011, I observed that West Eurasian populations were too close (measured by Fst) to allow for long periods of differentiation between them. By implication, there must have been a "common source" of ancestry uniting them, which I placed in a "womb of nations" of the Neolithic Near East. I proposed that migrations out of this core area homogenized West Eurasians, writing:

In Arabia, the migrants would have met aboriginal Arabians, similar to their next door-neighbors in East Africa, undergoing a subtle African shift (Southwest_Asians). In North Africa, they would have encountered denser populations during the favorable conditions of MIS 1, and by absorbing them they would became the Berbers (Northwest_Africans). Their migrations to the southeast brought them into the realm of Indian-leaning people, in the rich agricultural fields of the Mehrgarh and the now deserted oases of Bactria and Margiana. Across the Mediterranean and along the Atlantic facade of Europe, they would have encountered the Mesolithic populations of Europe, and through their blending became the early Neolithic inhabitants of the Mediterranean and Atlantic coasts of Europe (Mediterraneans). And, to the north, from either the Balkans, the Caucasus, or the trans-Caspian region, they would have met the last remaining Proto-Europeoid hunters of the continental zone, becoming the Northern Europeoids who once stretched all the way to the interior of Asia.

The new paper confirms the last two of these migrations. The remainder involve parts of the world from which no ancient DNA has been studied.

The first migration (early Neolithic) is already uncontroversial, but the paper includes data from Spanish early farmers that are also Sardinian- and LBK-like. The "Sardinian" Iceman was no fluke. It is now proven that not only the LBK but also the Spanish Neolithic came from the same expansion of Mediterranean populations which survives in Sardinia. The authors write:

Principal components analysis (PCA) of all ancient individuals along with 777 present-day West Eurasians4 (Fig. 2a, SI5) replicates the positioning of present-day Europeans between the Near East and European hunter-gatherers4,20, and the clustering of early farmers from across Europe with present day Sardinians3,4,27, suggesting that farming expansions across the Mediterranean to Spain and via the Danubian route to Hungary and Germany descended from a common stock.

The second migration went into eastern Europe:

The Yamnaya differ from the EHG by sharing fewer alleles with MA1 (|Z|=6.7) suggesting a dilution of ANE ancestry between 5,000-3,000 BCE on the European steppe. This was likely due to admixture of EHG with a population related to present-day Near Easterners, as the most negative f3-statistic in the Yamnaya (giving unambiguous evidence of admixture) is observed when we model them as a mixture of EHG and present-day Near Eastern populations like Armenians (Z = -6.3; SI7).

The EHG (Eastern European Hunter-Gatherers) are likely Proto-Europeoid foragers and the Yamnaya (a Bronze Age Kurgan culture) were a mixture of the EHG and something akin to Armenians.The "attraction" of later groups to the Near East is clear in the PCA: hunter-gatherers on the left side, the Near East (as grey dots) on the right side, and Neolithic/Bronze Age/modern Europeans in the middle. The second migration may very well be related to the Uruk expansion and the presence of gracile Mediterranoids and robust Proto-Europeoids in the Yamna:

The Yamna population generally belongs to the European race. It was tall (175.5cm), dolichocephalic, with broad faces of medium height. Among them there were, however, more robust elements with high and wide faces of the proto-Europoid type, and also more gracile individuals with narrow and high faces, probably reflecting contacts with the East Mediterranean type (Kurts 1984: 90).

The authors present a table of Fst values which confirms the homogenizing influence of migrations from the Near East. The WHG group has an Fst=0.086 with Armenians, but the LBK farmers have only 0.023. The EHG group has an Fst=0.067 with Armenians, but the Yamnaya steppe people have only 0.030. Someone might argue that it is the Armenians that are receiving genes from Europe, but the same pattern holds even for the Bedouins, for which admixture with Europeans seems far-fetched: 0.106 to 0.043 and 0.093 to 0.060. It is now clear that the "glue" that did not allow West Eurasian populations to drift very far apart were migrations from the Near East.

The (partial) demise of the farmers

It seems that the legacy of the early farmers suffered two hits, which is why only in Sardinia and (to a lesser degree) in southern Europe that they have persisted as the major component of ancestry. The first blow came during the Neolithic:

Middle Neolithic Europeans from Germany, Spain, Hungary, and Sweden from the period ~4,000-3,000 BCE are intermediate between the earlier farmers and the WHG, suggesting an increase of WHG ancestry throughout much of Europe.

And the coup de grâce after the 5kya mark:

We estimate that these two elements each contributed about half the ancestry each of the Yamnaya (SI6, SI9), explaining why the population turnover inferred using Yamnaya as a source is about twice as high compared to the undiluted EHG. The estimate of Yamnaya related ancestry in the Corded Ware is consistent when using either present populations or ancient Europeans as outgroups (SI9, SI10), and is 73.1 ± 2.2% when both sets are combined (SI10). [...] The magnitude of the population turnover that occurred becomes even more evident if one considers the fact that the steppe migrants may well have mixed with eastern European agriculturalists on their way to central Europe. Thus, we cannot exclude a scenario in which the Corded Ware arriving in today’s Germany had no ancestry at all from local populations.

Confirmation of the Bronze Age Indo-European invasion of Europe

In 2012 I had used the paltry data on a handful ancient DNA samples to observe that in ADMIXTURE modern Europeans had a West Asian genetic component (peaking in "Caucasus" and "Gedrosia") that pre-5kya Europeans didn't. I proposed that the Bronze Age migration of the Indo-Europeans spread this component:

But there is another component present in modern Europe, the West_Asian which is conspicuous in its absence in all the ancient samples so far. This component reaches its highest occurrence in the highlands of West Asia, from Anatolia and the Caucasus all the way to the Indian subcontinent. [...] Nonetheless, some of the legacy of the earliest Indo-European speakers does appear to persist down to the present day in the genomes of their linguistic descendants, and I predict that when we sample later (post 5-4kya) individuals we will finally find the West_Asian piece that is missing from the European puzzle.

This prediction is now confirmed:

This pattern is also seen in ADMIXTURE analysis (Fig. 2b, SI6), which implies that the Yamnaya have ancestry from populations related to the Caucasus and South Asia that is largely absent in 38 Early or Middle Neolithic farmers but present in all 25 Late Neolithic or Bronze Age individuals. This ancestry appears in Central Europe for the first time in our series with the Corded Ware around 2,500 BCE (SI6, Fig. 2b, Extended Data Fig. 1).

I was a little puzzled with the "Ancient North Eurasians" recently proposed as a "third ancestral population" for Europeans: it seemed to be a tertium quid that spread after 5kya, but very different geographically than the "West Asian" component. But:

These results can be explained if the new genetic material that arrived in Germany was a composite of two elements: EHG and a type of Near Eastern ancestry different from that which was introduced by early farmers (also suggested by PCA and ADMIXTURE; Fig. 2, SI5, SI6).

So, it seems that there is no contradiction after all and both EHG (which is related to "Ancient North Eurasians") and another type of Near Eastern ancestry (=West_Asian) arrived after 5kya.

1939 strikes back

It is amazing how well this was anticipated by Carleton Coon in 1939. Back then much of West Eurasia was an archaeological/anthropological terra incognita, there was no radiocarbon dating, no DNA, no computers, not even serious multivariate statistics. And yet:

We shall see, in our survey of prehistoric European racial movements, 8 that the Danubian agriculturalists of the Early Neolithic brought a food-producing economy into central Europe from the East. They perpetuated in the new European setting a physical type which was later supplanted in their original home. Several centuries later the Corded people, in the same way, came from southern Russia but there we first find them intermingled with other peoples, and the cul-tural factors which we think of as distinctively Corded are included in a larger cultural equipment. [...] On the basis of the physical evidence as well, it is likely that the Corded people came from somewhere north or east of the Black Sea. The fully Neolithic crania from southern Russia which we have just studied include such a type, also seen in the midst of Sergi's Kurgan aggregation. Until better evidence is produced from elsewhere, we are entitled to consider southern Russia the most likely way station from which the Corded people moved westward.

And in 2015:

Our results support a view of European pre-history punctuated by two major migrations: first, the arrival of first farmers during the Early Neolithic from the Near East, and second of Yamnaya pastoralists during the Late Neolithic from the steppe (Extended Data Fig. 5).

In 1939:

Linguistically, Indo-European is probably a relatively recent phenomenon, which arose after animals had been tamed and plants cultivated. The latest researches find it to be a derivative of an initially mixed language, whose principal elements were Uralic, called element A, and some undesignated element B which was probably one of the eastern Mediterranean or Caucasic languages. 5 The plants and animals on which the Somewhere in the plains of southern Russia or central Asia, the blending of languages took place which resulted in Indo-European speech. This product in turn spread and split, and was further differentiated by mixture with the languages of peoples upon whom it, in one form or other, was imposed. Some of the present Indo-European languages, in addition to these later accretions from non-Indo-European tongues, contain more of the A element than others, which contain more of the B. The unity of the original " Indo- Europeans," could not have been of long duration, if it was ever complete.

In 2015:

These results can be explained if the new genetic material that arrived in Germany was a composite of two elements: EHG and a type of Near Eastern ancestry different from that which was introduced by early farmers (also suggested by PCA and ADMIXTURE; Fig. 2, SI5, SI6). We estimate that these two elements each contributed about half the ancestry each of the Yamnaya (SI6, SI9), explaining why the population turnover inferred using Yamnaya as a source is about twice as high compared to the undiluted EHG.

The EHG is still flimsy as it's only two individuals from Karelia and Samara who are very similar to each other. It's hard not to imagine that the hunter-gatherer from Russian Karelia (outside any proposed PIE homeland) would be speaking a similar language as his Samara counterpart. Did they both speak "element A" and was PIE formed when the "southern" steppe hunter-gatherers came into contact with "element B" people from the Caucasus? Short of a time machine, we can never say for sure. This might very well be an answer to the conundrum of Uralic/Proto-Kartvelian borrowings. There is simply no geographical locale in which these two language families neighbor each other: Northwest, Northeast Caucasian speakers and the pesky Greater Caucasus intervene. But, maybe there was no such locale, and these borrowings aren't due to some "PIE people" living adjacent to Uralic and Proto-Karvelian speakers but the "PIE people" being a mix of an element A (EHG) that was (or interacted with) Uralic and another element B (Armenian-like) that was (or interacted with) Proto-Kartvelian.

Urheimat (or not?)
The authors of the current paper are agnostic about the PIE homeland:

We caution that the location of the Proto-Indo-European9,27,29,30 homeland that also gave rise to the Indo-European languages of Asia, as well as the Indo-European languages of southeastern Europe, cannot be determined from the data reported here (SI11). Studying the mixture in the Yamnaya themselves, and understanding the genetic relationships among a broader set of ancient and present-day Indo-European speakers, may lead to new insight about the shared homeland.

Whatever the ultimate answer will be, it seems that Coon was right that "The unity of the original " Indo- Europeans," could not have been of long duration, if it was ever complete." If PIE=EHG (as Anthony and Ringe suggest), then "from the crib", PIE got half its ancestry from a non-IE, Near Eastern source. Conversely, if PIE=Near East (as I suggested) then "from the crib", PIE got half of its ancestry from a non-IE, Eastern European source. The "Yamnaya" seems to max out in Norwegians at around half, which means that they are about a quarter Proto-Indo-European genetically, regardless of which theory is right.
These two possibilities (as well as the third one of PIE being neither-nor, but rather a linguistic mixture of the languages of the EHG and Near East) are testable. The Anthony/Ringe version of the steppe hypothesis predicts pre-Yamnaya expansions from the steppe. Whether these happened and what was their makeup can be tested: if they did occur and they did lack "Near Eastern" ancestry, then the steppe hypothesis will be proven. PIE in the Near East, on the other hand, predicts that some PIE languages (certainly the Anatolian ones) will be a "within the Near East" expansion. If such migrations did occur and they lacked "EHG" ancestry, then some variant of the Gamkrelidze/Ivanov model will be proven. Or, the truth might be that everywhere where Indo-Europeans arrive they carry a blend of "West Asian" and "EHG", supporting the third possibility. Time will tell.

In the interim, I am curious about how much Yamnaya ancestry existed in different parts of Europe (all of the post-5kya samples in this study come from Germany, with a couple from Hungary). In northern Europe, all populations seem to have less Yamnaya ancestry than the Corded Ware: there it must have declined. But, modern Hungarians have more than Bronze Age Hungarians: there it must have increased.Germany and a slice of Hungary is a very narrow window through which to see the whole of Europe and these results must be tested by looking at samples from beyond the "heartland". I do hope that some kind of Moore's law operates in the world of ancient DNA, and in three more years we'll be reading studies about thousands of ancient individuals.

bioRxiv doi: http://dx.doi.org/10.1101/013433Massive migration from the steppe is a source for Indo-European languages in Europe

We generated genome-wide data from 69 Europeans who lived between 8,000-3,000 years ago by enriching ancient DNA libraries for a target set of almost four hundred thousand polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies and to obtain new insights about the past. We show that the populations of western and far eastern Europe followed opposite trajectories between 8,000-5,000 years ago. At the beginning of the Neolithic period in Europe, ~8,000-7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary, and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ~24,000 year old Siberian6. By ~6,000-5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ~4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ~3/4 of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ~3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for the theory of a steppe origin of at least some of the Indo-European languages of Europe.

Great analysis, Dienekes. Put that EHG derived from WHG of the Southern European Refugia and you'll understand where was the origin of R1a* and R1b*. About the mixture of Uralic and Caucasian we'll se next if it happened in South Russia or nearby the Alpine zone (Caucasian languages were spoken also in Western Europe: Sardinian and Basque).

Firstly, I find it mind-boggling that the seafarer-agriculturalists who settled Cyprus, Crete, the Greek Islands, and much of the Mediterranean are supposedly identical to the North Anatolian/ Southern Balkan people who became LBK. But perhaps they weren't: there is definitely a pretty large PC2 gradient within the Early Neolithic population (~3 times as much as the PC1 variation caused by WHG/W Asian agriculturalists admixture!).

Secondly, in much of N Europe, say around and west of the Elbe river, Corded Ware from the archaeological perspective is not intrusive, but a parallel occurrence to late TRB with an eventual merger. That brings up two questions: (i) can this be identified in an East-West gradient of "Yamnaya admixture," and (ii) is it possible that LBK --> TRB already spoke a language related to pre-PIE. One should note that LBK arrived in the Ukraine via the Elbe-Saale region and Poland, from the NW. So, whatever language hybridization took place in the Yamnaya region might have a predecessor during the Eastern LBK migrations (West Asian + EHG).

It really bothers me that no Neolithic farmer has been found who is not some sort of half breed. There must have been many Near Eastern farmers living in Europe for hundreds and thousands of years before admixing with hunter/gatherers. Where are their remains, their dna? It is important to me to know what those original farmers were like and how they compare with modern and ancient Near Easterners. I do not like the use of modern and admixed proxies like the Bedouin B and the Nganasans.

My personal opinion is that these type of deductions about the ancestry of Europeans will have to be completely revised or thrown out when more ancient dna is tested especially of unmixed first Neolithic farmers, and ancient remains from the South of Europe, North Africa, the Near East and Cyprus.

'We caution that the sampled Yamnaya individuals from Samara might not be directly ancestral to Corded Ware individuals from Germany. It is possible that a more western Yamnaya population, or an earlier (pre-Yamnaya) steppe population may have migrated into central Europe, and future work may uncover more missing links in the chain of transmission of steppe ancestry.'Apparently, the status of Yamnaya is not resolved at all. If they weren't the direct ancestors of corded Ware, and too R1b M269-y to be the ancestors of Indo-Iranians, it is still most likely they became extinct. EHG roots and other genetic evidence also suggest the Yamnaya presence in the Pontic Steppes was not continuous to previous cultures, such as Sredni Stog. IMO, Yamnaya is now established as nothing but a close and possible unfortunate cousin to Corded Ware, not its ancestor!

Thank you Dienekes! That was an incredibly intelligent and insightful analysis! The terrible force of the expansion of that Yamnaya like ancestry may lie in the new inventions that kept flowing from the east to the west and from the south to the north. IE languages were a part of that process but they were not the only language family spreading the new ways of life to their surroundings and we definitely should reconsider the position of the Basque language, Northwest and Northeast Caucasian languages, Kartvelian languages and Uralic languages in this Bronze Age upheaval and the impact it exerted on whole Europe.

"Someone might argue that it is the Armenians that are receiving genes from Europe, but the same pattern holds even for the Bedouins, for which admixture with Europeans seems far-fetched: 0.106 to 0.043 and 0.093 to 0.060. It is now clear that the "glue" that did not allow West Eurasian populations to drift very far apart were migrations from the Near East."

Yes, that's certainly one explanation but, since the Near Eastern component in Yamnaya is not the same Near Eastern component as came to Europe with farmers, the other option is worth serious consideration. Remember what Raghavan et al. wrote about ANE: "Thus, if the gene flow direction was from Native Americans into western Eurasians it would have had to spread subsequently to European, Middle Eastern, south Asian and central Asian populations, including MA-1 before 24,000 years ago." If we find ANE-rich Yamnaya ancestry in Bedouins it fits with the presence of ANE ancestry in the Middle East (and likely in Bedouins).

Notice also that Fst is the highest between Amerindians and all and any Old World populations.

Thanks for the description of physical appearance of the Yamnaya population. But the new study also shed some light on R1a (for example this Z280 from the Lusatian Culture site in eastern Germany, as well as this Mesolithic Karelian hunter). I've found a reconstruction by M.M. Gerasimov of that Mesolithic R1a male (hunter-gatherer who lived 5500 years ago) from Karelia (Lake Onega island):

"The second migration may very well be related to the Uruk expansion..."

That is certainly one possibility. Another is the earlier Northern Ubaid expansion. If I'm remembering correctly, the earliest kurgans appear south of the Caucasus, in a culture influenced by the Northern Ubaid, rather than Uruk.

Yet another possibility is an expansion of pastoralists northward along the eastern shore and east of the Caspian. I don't think some combination of the above out of the question, either.

In Mehrgarh I, we find a fully developed, pre-pottery Neolithic village, with clear parallels in the Kurdish Zagros. If sited in the Levant, it would be PPNB, which itself is suggested to be a northern influence in that region. The interesting suggestion is that, by the eighth millennium BCE, there was a southern and eastern expansion out of more-or-less this same area from which we're proposing a northern expansion some millennia later.

Dienekes- I wouldn't be too confident in praising the physical anthropologists of old too much. Eg there's no 'great Mediterranean race" from Atlantic to "Classic Mediterranean" to East Med type; nor are the "Dinarics" (modern balkans) related to "Armenioid race "

Va-Highlander I think those Georgian kurgans require further , independent datingWhatever the case , scholars have now been lookng toward Central Asia (Zargos) as the "catalyst" for Majkop, rather than Mesopotamia.

@ Kristiina

"Terrible forces" and Bronze Age "Upheavals" ? Let's not turn this into a Hollywood blockbuster .

@ Rokus Whilst we still need to sample more western Yamnaya areas ; you could of course be right . And despite the early presence of R1b in Sammara, the possibility remains that the M4-3 R1b arrived from west-central asia. I also think the extinction of yamnaya genomes are a recent phenomenon - the slavic expansions, etc. The surprising world of aDNA might prove me wrong, of course.

'Secondly, in much of N Europe, say around and west of the Elbe river, Corded Ware from the archaeological perspective is not intrusive, but a parallel occurrence to late TRB with an eventual merger.'Indeed, and possibly the westernmost expanse of 'Corded Ware' being attested well within the TRB megalith horizon of the Netherlands was initially as utterly unaffected by the Yamnaya component as Baalberge and Bell Beaker were. Apparently, the earliest YDNA R in western Europe was not necessarily accompanied by ANE. Early Neolithic Troc3 in Spain attested R1b1 already long before any Yamnaya-emigration ever hypothesized: 5178-5066 cal BCE! Also the circumstance that the sampled Yamnaya turned out already differentiated towards the Balkan-Anatolian-Armenian subset of R1b1 (Z2103) implies a much wider network of IE cultures in existence already at an early stage - not unlike the broader homeland concept that Malloy introduced, albeit as 'strawman' to give credence to his now apparently obsolete idea of a Sredny Stog source population. What we see now is just one stage in the process of intermingling throughout the network.

@Arch Hades (whose original comment is in the "genetic map of the British population" post, but was probably intended for this one)

As per the "Bronze Age Indo-European invasion..." post, I think it's clear that I do not advocate an early (Anatolian or otherwise), but a late expansion of Indo-European languages.

I don't really have an opinion on how far back the split of Anatolian and the rest of PIE can be put. That whole literature is somewhat opaque, and the "wheeled vocabulary" argument is not informative about PIE itself. The first-order split of PIE must have happened in Neolithic times, which isn't really controversial whether one thinks it's migration from the steppe into the Balkans or "within the Near East".

If the early farmers of Europe (who came from Anatolia according archaeologists) lacked the "West Asian" component, then the Anatolian farmers must have lacked it too. So, I think that the "West Asian" component spread into Anatolia from the east, which might also be an explanation for the physical anthropological change that takes place in Anatolia in the Bronze Age. So, my best bet is that the "West Asian" component is coming from the east, from what I call "highland West Asia", which includes areas known as "eastern Anatolia", "northern Mesopotamia", "Armenian highlands", "Transcaucasia", and so on. Localizing it any further is hopeless at this point. The one constraint is to be close to Proto-Kartvelians (wherever those may have been in Neolithic times).

Thanks Dienekes, I'll just repost my question in here which you just answered, lol.

IMO, and I think this is what Dienekes is trying to say.... the PIE people are either The 50/50 mixture of the pseudo Armenian and EHG/Karelians that formed the Yamna culture if we follow the steppe hypothesis. Or just the pseudo Armenians themselves if we follow Armenian highland hypothesis.

There's no major linguistic hypothesis of the PIEs being the Karelians or in the vicinity of that area.

I take it you've abandoned Neolithic Anatolia, Dienekes? I don't see how Neolithic Anatolia would be strengthened by these turn of events. The ancient Sardinian language was non IE and The Near Eastern element that influenced the Steppe was Armenian like, not Sardinian like, like all the earliest European farmers in Hungary/N Italy/Germany/Spain.

Since there was major genetic turnover Central/Northern and Western Europe after the Neolithic it's likely the Steppe expansions brought IE to those areas and replaced the non IE element that those farmers spoke, other than the old Paleo Sardinian language.

How could one fit Neolithic Anatolia into a larger frame with this new information?

What is your opinion of the possibility [for at least some] of the "pre Greek" Neolithic people of the Aegean and Crete being Indo-European and speaking perhaps a language similar to Anatolian? There are some linguists like V. I. Georgiev who believed this to be the case, and even Mallory 1989 notes that there is a non Greek but fully Indo-European substrate in Greek [along with a large non IE substrate]

Is that impossible via your model of "West Asian" and "Bronze Age Indo-European invasion" of Europe?

BTW, isnt the R1b found in this Yamna group here of the Greek-Balkan-West Asian variety? What do you make of that?

Nice paper. Best evidence of the mid- Neolithic Northern migration into Europe that I have ever seen. Yamna seems to be the most likely source if not the only source.

It is also the nicest summary I have ever seen of the huge holes in our knowledge and the twisting that is being done to make things fit.

The only truly pre-neolithic samples are the Pleistocene Hunter Gathererss (43-22 BCE) way off to the East. In K=16 they bear almost no resemblance to the other samples.

The neolithic commenced 10 BCE in the East, and arrived in Europe 5-7 BCE). ALL the other genomes that we know about are fully in the neolithic era (6 BCE or younger) or more recent.

There has been a tendency in the past to categorize samples by what fits the conservative message, rather than reality. So samples of the same age may be classed as hunter gather or farming on very little evidence. I have argued about this previously. For example Iceman is classed as a farmer although he was actually living a hunter gatherer lifestyle. This makes me suspicious of the allocations of genomes to groups.

There are two very divergent groups labelled hunter gatherers. The Northwestern Europeans and the Western Europeans. I was surprised at how separate they were genetically from each other. THERE ARE NO SOUTHERN EUROPEAN HUNTER GATHERER SAMPLES, even though due to climate this would have been the most numerous group prior to the neolithic.

WE HAVE NO IDEA WHAT SOUTHERN PRE NEOLITHIC HUNTER GATHERERS LOOKED LIKE GENETICALLY. If you class Iceman (Oetzi) as a representative of the pre-neolithic hunter gatherers, then they were indistinguishable from the bulk of the neolithic samples. This is what has been done for the Neolithic Swedish and La Brana samples in similar circumstances, but Oetzi is different, because that upsets the required story of Near Eastern replacement. Twisting.

From this paper it looks like an old Eurasian (Gravettian? Aurignacian?) hunter gatherer population contributed to Gravettian age MA-1. Descendants of the this population were still travelling around Europe in the Neolithic (NW and Western Hunter Gatherers). These northern hunter gatherers are the third component in most European countries but are largely absent from Southern Europe, suggesting this was not part of their range. So who lived in Southern Europe before the neolithic?

Modern Europeans seem to be about half the southern component (Orange), half the northern component (Green) with variable amounts of the Northern Hunter Gatherer (blue). There is a rough gradient of Southern/Northern from South West to North East, and a a gradient of Northern Hunter Gatherer from North to South.

At this stage, from this paper, the Northern component (Green) seems to have arrived in the mid Neolithic some time AFTER the LKB pottery era. Yamna-type herders seems reasonable.

We dont know when the Southern component (Orange) arrived. All we know is that it was present in the early Neolithic from Spain to Stuttgard. it could have been there since the Aurignacian. Arrival from the near East in the neolithic is pure assumption.

We dont even know when the Northern Hunter Gatherers (Blue) arrived. They could have arrived in the Aurignacian, Gravettian or even been Johnny-come-lately nomadic folk who arrived from northern Asia in the mid Neolithic. Or all of the above (multiple inputs over a long period of time). All we know is that they were present in the Neolithic.

IMO there has been no convincing link between the southern component and neolithic era population replacement from the Near East. This component is more likely in my opinion to be initially a post LGM expansion of mediterranean hunter gatherers (climate change), followed by neolithic expansion from the south to the north (adoption of farming technology).

The absence of WHG in Spain and Tuscany in your figure 3 seems suggestive.

Yamna(/-like) presumably spread there, and at some point mixed with an ENF population, without mixing on the way with populations in other areas, like CW, that have WHG?

Is that indicative of a fairly rapid expansion of Yamna-like right across the continent or of multiple waves?

Or are Spain and Tuscany entirely a product of some movement of (ENF+Yam) that totally replaced the previous population?

Like you said, we need more ancient samples, preferably thousands, from across Europe to build a better picture of ancient migrations.

(Bodmer et al seem pretty useless, hogging all the British samples for some sketchy PC establishment paper. Where and when did Yamna in Britain come from Bodmer? How about sharing the samples and letting the scientist do some serious work on them? I am waiting for Bodmer to tell us something stupid like "we all come from Africa and mass immigration is right on!")

Coon's account of the genesis of the Indo-European languages was based on Tubetskoy's then very novel (and still controversial) theory that involved contact between a Central Asian and a Caucasic language.

Could I make a request from the great minds of this board? Could someone write up a quick chart for us? It's hard to keep up with the latest theories and findings.

What I mean is something like this:

There are n major components to European ancestry. (Is it 3 or 4?)

The 3 (4?) components are referred to as (insert acronyms).

The acronyms stand for...

The modern ethnic group retaining the largest number of XYZ alleles is the...

I can't promise you I'd buy you a beer from afar, but you will be my hero.

Also, did I read this correctly that people are postulating that Norwegians -- who ALL previous studies indicate have very little "eastern" blood, and who do not look like eastern populations (old or new) (at all) are supposedly the group retaining the most Yamnaya genes? That just cannot be right.

I don't really agree with ur interpretation First of all; the R1b samples from yamnaya are not ancestral to weatern european ones Secondly ; I'm not sure I'm wholly convinced the shift to ANE in Central Europe should be laid at the doorstep of yamnaya alone .

"So, my best bet is that the "West Asian" component is coming from the east, from what I call "highland West Asia", which includes areas known as "eastern Anatolia", "northern Mesopotamia", "Armenian highlands", "Transcaucasia", and so on."

Or maybe even further east than that.

.

@Annie Mouse

"We dont know when the Southern component (Orange) arrived. All we know is that it was present in the early Neolithic from Spain to Stuttgard. it could have been there since the Aurignacian. Arrival from the near East in the neolithic is pure assumption."

If you had orange HGs over a wide range and one group developed agriculture and expanded then you could have orange farmers moving into the territory of orange HGs with no way of knowing.

Mike, I wrote what I wrote because I was shocked to see that there was no old WHG in modern populations in Extended Data Figure 1 K=20. It has completely disappeared. For example Motala is 100% of old WHG and modern Norwegians have none of it. This is probably not the whole story and there are technical issues that should be discussed. By contrast, Figure 3, p. 23 is much more balanced and may be closer to the truth. According to it, Norwegians have c. 16% of old WHG and Estonians almost 40%. On the basis of this, I understand that La Brana should be most of all Estonian like.

Upheaval may also be due climatic changes, as I noted earlier on another blog.

@Anne MouseAnne, I don't know if you are aware that what you say is in line with my theory of the Italian Refugium (of many hgs, both Y and mt, but in this case of R1b1-L389* and subclades). Why the conclusions of this paper are wrong? Because, very likely, not having found in Yamnaya any R-L51 (which is centered in Italy, in spite of all what many have said in these days, and like the first map done by Argiedude and me demonstrated many years ago), demonstrates that R-L51 and subclades weren't there but very likely in Western Europe, from Italy to Iberia 7500 years ago, and from Iberia to Ireland (I have written a lot about that and about the link of the oldest R-L51 subclade CTS6889- between Tuscany and Ireland).Here are the ages: YBP BCE SUM SNPs in Block Chr Y HG8500 6500 91 4 R1b1a2a1 - L51and not having found R-L51 or subclades in Yamnaya does mean very likely that it hasn't ever been there.Also about R-L23 it isn't said that the origin is there. I am R-L23/Z2110 and have 16 SNPs in common with a Basque and with the Englishman Barnum/Winterowd. After I have 66 private SNPs from my Full Genome.And also about R-M269 I have some doubts that the origin is there: my acquired cousin Fabrizio Federighi is the alone to have the mutations PF7566 and PF7569 beyond the other known and finds so far a link with a Greek. These data are all in the smal's tree in the "R1b1a2 (P312- U106-) DNA Project" at FTDNA.

"WE HAVE NO IDEA WHAT SOUTHERN PRE NEOLITHIC HUNTER GATHERERS LOOKED LIKE GENETICALLY. If you class Iceman (Oetzi) as a representative of the pre-neolithic hunter gatherers, then they were indistinguishable from the bulk of the neolithic samples."

Annie Mouse,

Ötzi was found in a region that can easily be reached from the NW Balkans along several rivers (even via the Danube). I have previously argued that the ~Ice Age Maximum (LGM) Balkans were geographically and demographically decoupled from the rest of Europe (likely with the exception of habitable zones in Italy - especially the-then extended Po valley); but, because the Bosporus was a land bridge and because of known seafaring skills, they were very much connected with NW Anatolia. So, ~45-25 kya, they were all the same (old Aurignacian+Gravettian WHG with a cline to the Near East), and then you have a 15 ky gap to the beginning of the Neolithic. Perhaps this explains why the early Neolithic samples are so incredibly narrow in PC1 - that takes more than a few generations, especially in a large, quickly growing, and geographically widely expanding population. But in PC2 they are rather far apart, meaning that (as today) they were sorted by latitude during LGM and only thrown into similar heaps during the westward Neolithic expansion.

I was thinking of the Leyla-Tepe culture, of Azerbaijan. It should also be noted that the Northern Ubaid and the Ubaid period of southern Mesopotamia are not necessarily the same cultural phenomenon.

"Whatever the case , scholars have now been lookng toward Central Asia (Zargos) as the "catalyst" for Majkop, rather than Mesopotamia."

I understand material goods point to northern Iran and southern Turkmenistan, yes, but I remain a bit unsure just what this is telling us. As you say, though, in any case the center-periphery model, with cultural and technological influence radiating outward from southern Mesopotamia and the Levant, seems no longer operative.

Given what we now know about ancient networks of trade and influence in this region, I don't think it's particularly difficult to push all the various PIE descendants into their proper locales at the proper times. Anthony's recent paper is something of a straw man, really. An early out-of-Anatolia model is, as he never tires of demonstrating, even more unlikely than his Kurgan model.

@Grey"If you had orange HGs over a wide range and one group developed agriculture and expanded then you could have orange farmers moving into the territory of orange HGs with no way of knowing."

Agreed.

@FantyThe line between the southern folk and the northern folk seems to cut across northern Spain with La Brana on the northern side. This is so even for different types of neanderthal ranges. Not sure why but I think it is to do with suitable game areas.

The main difference between EEF and the modern "West Asian" component is.

Both obviously are by majority Neolithic proto farmer with the only difference.

West Asian(proto pastoralists) being Proto-Farmer(~70%)+ ANE(~30%).

While EEF is ENF(80%)+ WHG(20%)

This could have happened in the Balkans but already in Western Asia, because especially Levantine and Anatolian populations have weirdly "significant" percentage of WHG like component, which could be the remains of EEF. And this component might have been altered by incoming ANE admixture in the east, which was the cause for the "West Asian" component. And incoming East African admixture in the South and Southwest which in turn caused the appearance of this "Southwest Asian" component.

And we do have South European mesolithic data (La Brana).

Now back to topic. I am still in favor of a Zagros/Alborz-North Mesopotamia origin of PIE.

I wouldn't rule out an Anatolian genetic influence in Europe. You even see juuuust a hint of it in the Italian sample. I'm running with an influx of J2a-M530 on my next video series. Variance distribution of that tribe in the Balkans and Southern Italy shows a perfect match with the movement of Neolithic technology into Europe as documented archaeologically.

The M67's and M92's are definitely Bronze Age immigrants. I'd bet the farm on that. I'm also 100% those "Mycenaean colonies" we start seeing in Italy after 1700 BC are Cretan. That would be when those latter groups started filtering in. And after 1300 BC - for reasons I'll cover in my next series - they started POURING in.

"Given what we now know about ancient networks of trade and influence in this region, I don't think it's particularly difficult to push all the various PIE descendants into their proper locales at the proper times. Anthony's recent paper is something of a straw man, really. An early out-of-Anatolia model is, as he never tires of demonstrating, even more unlikely than his Kurgan model"

A Neolithic dating is out of the question, linguistically. But as you point out, the Kurgan models is like a house of cards...

Whatever the case, we see events (c. 3500 BC) linking broad regions of west Eurasia: the Copper Age Balkan collapse coincides with the rise of Majkop, which in turn has clear southern 'impulses', but yet difficult to define as to exact nature. We are seeing a triangle (? perhaps a trapezoid) linking the Carpatho-Danube region, Caucasus, central Asia, and Anatolia-Aegean. PIE could have first been spoken anywhere here.

P.S. If the theory of the origin of the IE languages from Yamnaya were true, I should be proud, because, being R-L23-Z2110, I would descend from those people who carried those languages to Western Europe (of course the R-L51 and subclades lived in Western Europe before, without finding this subclade in Eastern Europe, but it is at least 8000 years old and not less than 5000 for descending from Yamnaya people), but also my line is in Western Europe from the same old time: link with a Basque and an Englishman 80 SNPs ago.

"Adding any Early or Middle Neolithic farmer results in EHG related genetic input into Late Neolithic populations being a poor fit to the data (SI9); thus, Late Neolithic populations have ancestry that cannot be explained by a mixture of EHG and Middle Neolithic."

You surely have to add a W Asian component, in addition, to explain Corded Ware, that way. I don't see that the data are inconsistent with West Pontic (e.g., ~Globular Amphora) similar to the middle-Neolithic Hungarian sample - but admixed with Bronze-Age Near-Eastern, plus EHG. Thus, the data might be explained by a W Pontic rather than steppe expansion, as well.

in R1a1a and Subclades Y-DNA Project there's five men belonged at the same subclade of korelian man, an Italian, a Tunisian, two belarusian and a Rusian, they are all R-M459, R1a1 * -M17- M198-, this branche is now called YP1272 sister of M198 subclade

The paper states they originate from yamnya and moved into Europe from Yamnya. Be these markers east-asian, siberian, near-eastern or whatever is irrelevant, they came from yamnya after settling in yamnya from elsewhere for an extended period.Clearly for the paper to claim this, means isotopes where done on the skeletral remains found in Europe by these yamnya people.

" I have previously argued that the ~Ice Age Maximum (LGM) Balkans were geographically and demographically decoupled from the rest of Europe (likely with the exception of habitable zones in Italy - especially the-then extended Po valley); "

There is no reason to assume that the Balkans were completely decoupled from Europe during the LGM. (Italy? Romania? Austria? Ukraine?)

I've seen you make many comments like this. I think it just reflects your antipathy for people from the Balkans.

" So, ~45-25 kya, they were all the same (old Aurignacian+Gravettian WHG with a cline to the Near East), and then you have a 15 ky gap to the beginning of the Neolithic."

There is no reason to assume there is a 15 ky gap in gene flow in Europe during the LGM.

There is no reason to assume that a complete merge between "Aurignacian+Gravettian WHG" related cultures during the Upper Paleolithic or Mesolthic.

None of these statements are supportable or even the most likely scenario.

Magnificent paper. I wouldn't hesitate to nominate this team for the nobel prize!

Although the authors remain cautious about the IE question, to me the evidence is quite clear-cut: I think the IE homeland was on the PC steppe (for R1b peoples) and in the forest zone to the north of it (for R1a peoples). Autosomally they were very similar, but they had different yDNA signatures.

A lot depends on the origin of R1b1a-P297 in Yamnaya. The possibility has been raised that this was from the West Asian element in Yamnaya. Its predecessor R1b1 was present in an Iberian EEF, in an EHG from Samara, and quite possibly it was also present at an early date in West Asia, since R1b1b and R1b1c are typically West Asian. But I think the most obvious explanation is that R1b1a-P297 in Samara was descended from the R1b1 in Samara. Same place, and just one mutation away. Also, if the IE element had arrived on the steppe from West Asia, linked with R1b1a-P297, the Corded people should have R1b1a-P297 as well, otherwise they couldn't be IE too. The presence of R1a1 in a Karelian EHG pretty much refutes the recently proposed hypothesis that R1a originated in West Asia. The geographical distance alone makes this unlikely, but it's even more unlikely since that EHG has no detectable West Asian autosomal admixture. And so far no Corded male carrying R1b has been found. On the other hand, pre-proto-IE might have developped among EHG groups who were in mutual contact, then there wouldn't have to be one haplogroup dominating the other.

Some have suggested that the predominant R1b1a2a2 in Yamnaya came from West Asia. But its direct ancestor R1b1a2a was also present in Yamnaya, as was R1b1a. So the only unattested link is R1b1a2, which may easily have arised from the local R1b1a. Also I note that they don't have the typical West Asian R1b-L277, but its ancestor. And this is exactly what would be expected if IE languages in West Asia were derived from the steppe. Also it has been criticized that this Yamnaya sample doesn't have R1b-L51, which is typical for Europe. But it does have its direct ancestor, R1b-L23. Quite possibly L51 didn't yet exist back then, but it was just a mutation away.

So, the impression being that Yamnaya was paternally EHG-dominated, it's likely that the pre-proto-IE language came from the EHG, because the PIE were a patriarchalic society.

I would derive Italo-Celtic from the Yamnaya in the Carpathian basin. Like Marija Gimbutas suggested, the Bell Beaker eastern group was an amalgam of Yamnaya and Vucedol influences. Vucedol was related to Baden, i.e. CO1-like EEFs. Italo-Celtic spread further along the Danube, but presumably the first Italic wave to Italy came directly from the Carpathian Basin and the western Balkans. Oscan-Umbrian however, which shares the *kw -> p innovation with some Celtic languages, probably arrived later, with the Protovillanova-wave from Bavaria and Austria.

Perhaps the typical Dinaroid cranial type of many Bell Beaker groups is merely the result of a mixture of a Yamnaya derived mesocephalic type with West Asian facial features and a local brachycephalic type related with the Danish brachycephals. As Coon wrote: „So high is the mesocephalic ratio, and except for Hungary, so infrequent the truly long-headed crania associated with this type, that the mesocephals are clearly one branch of the main type, and not the product of local mixture with long heads. Morphologically, the mesocephals are essentially Bell Beaker.“ I think the EEF rarely had prominent, convex noses, that's probably associated with the later West Asian admixture carried by the IE. Gabriel Byrne looks like a typical example, I think his low interorbital distance is also typically West Asian.

As for the Corded Ware, we have to distinguish between their burial rite and their material culture. The material culture has a clear east -> west temporal gradient. In Poland it was already present before 2900 BC, while in the west mostly after 2700 BC. The burial rite on the other hand seems to have partly earlier, local roots and doesn't seem to be exclusively linked to the spread of EHG admixed people, although the latter surely contributed to its final success. I used to think that the Fatyanovo group in Russia was derived from the central European Corded Ware. But according to David Anthony, Fatyanovo is clearly older. Interestingly, the Fatyanovo cranial remains cluster with central European Corded people, suggestive of a biological link. The Fatyanovo pottery seems to unite Corded Ware and Globular Amphora traits, which might mean that they were some kind of basal outgroup of both cultures. According to wikipedia, Fatyanovo may be derived from the Middle Dneper group, which is roughly in the geographical midpoint of all Corded Ware groups. The Corded Ware was undoubtedly at the root of Balto-Slavic, but it's quite clear that it also had an important effect on the formation of Germanic, which to me looks like an amalgam of pre-proto-Italo-Celtic and pre-proto-Balto-Slavic. At this early stage, the different IE variants were perhaps still similar enough to enable a fusion.

Congratulations to Eupedia. Somehow they managed to guess it right all the time.

I would like to mention one other concern and reiterate another result that no one has commented on.

Firstly, Fig. 2a was created using all (ancient and modern) data, with the latter almost outnumbering the former by an order of magnitude. I think it is unfortunate that we don't have a similar figure just based on the ancient data, with the modern ones projected. Modern West Eurasian DNA has new contributions from historic or otherwise relatively recent influxes from Central Asia, East Asia, Africa, and even South Asia - all of which distract and pollute the data.

Both Figs. 2a and S5.2 show a very tight grouping of the Early Neolithic along PC1 (and also along the diagonal that is perpendicular to the West Asian population cline). In both cases, The Spanish samples are to the bottom-left, perhaps indicating their Cardial Pottery distinction from LBK as arising from a more southern (N Levantine vs. N Anatolian/ Balkan) origin, and also having a more southern HG admixture (S/SW Mediterranean vs. C Europe). But, more importantly, you don't see this type of tight grouping in any known modern populations that are admixed from predominantly two, distant groups. Modern African Americans fit a wide cline between Europeans and Africans, and so do US Americans of Mexican origin, in terms of Natives and European points of origin. The tight clustering of the Early Neolithic population indicates to me that they mostly have their own, separate origin - namely, in N Anatolia and the Balkans.

By the way, I'm not so sure if it's appropriate to call the Yamnaya population "unfortunate" if they were, as it seems, part of the PIE and fathered more offspring in Europe than any other archeological group...

The 5600 BC Samara forager's R1b was R1b1(xR1b1a1-M73, R1b1a2-M269). So we really have no idea what kind of R1b he was. However, if the new Y-Full coalescence dates are correct, which put M269 at 12200 (+- 2000) years old, 3-7000 years earlier, then the Samara forager cannot be closely related to the main European line at all.

The Yamnaya samples were mostly R1b1a2a2-Z2103, with one R1b1a2*-L23(xR1b1a2a1-L51, R1b1a2a2-Z2103), and one man who was derived for P297 and ancestral for L51, but whose L23 and Z2103 status is unknown. So there is no reason to suppose he is actually P297*; he may well be L23/Z2103 like the rest.

The 5500 BC Karelian forager is R1a1(xR1a1a-M198). Again according to Y-Full's tree, M198 coalesced 8500 (+- 1300) years ago, so probably this guy is not on the main line of R1a evolution either.

The Corded Ware guys have R1a1a1-M417* - Z282 doesn't show up until the Late Bronze Age. With the very few samples we have, we cannot argue from the absence of any haplogroup, only from their presence.

I can't agree that foragers carrying R1 without farmer admixture is evidence against a West Asian origin of R1b-L23 and R1a-M417. The shared ancestry between West Asia and Russia could long predate the arrival of Neolithic farmers. In any case, since we have an R1b1 Spanish farmer from 5000 BC in this paper, we know for sure that R1b was already found far beyond the steppe and in people with distinct autosomal types.

'By the way, I'm not so sure if it's appropriate to call the Yamnaya population "unfortunate" if they were, as it seems, part of the PIE and fathered more offspring in Europe than any other archeological group...'How, if Yamnaya is just part of PIE, like hypothesized in the broader homeland alternative once ridiculized by Mallory. Probably it was on the receiving end from an older source culture. There is no guarantee whatsoever that PIE truly was a Yamnaya affair. The puzzle is far from complete. Did Yamnaya R1b1a2a2 samples derive from the same people that expanded already before into the Balkan at 4200 BC? This subclade survived in Anatolia and Balkan that may have been populated by this event, what means that 3300 calBC Yamnaya was nothing but a late descendent AND a dead end. Without Yamnaya as the exclusive PIE glue it becomes tenuous to consider a steppe origin at all. Attention is already shifting to the eastern Corded Ware horizon for a more complete picture, then where will this bring us? Away from the steppes for sure, in both time and space. Fatyanovo has a more obvious link to Funnel Beaker and Pitted Wares, what may reduce the Dnieper groups to a transitional role that ultimately derive from Funnel Beaker, what should come as no surprise for those that believe the wheel was a PIE invention.

@ Annie " doesnt have to be the Yamnaya. Just someone similar autosomally."

Ok Annie; yes I agree it could be *anyone * autosomally similar to Yamnaya which moved west to Germany to replace the LN farmers there.

But, whoever the people which colonized Late Neolithic Germany were, they were not of the R1b branch (z-2103) as the currently sampled Yamnayans. They are thus not one and the same "causal' population, but merely an autosomally related, R1a predominant one, which had otherwise diverged several thousand years earlier.

So then, if the sampled Samaran Yamnayans neither invaded eastern Europe, nor western Europe, I fail to see how this study "proves" the Kurgan hypothesis just becuase there was a large 'eastern shift' in the autosomal profile of central European Bronze Age groups ?

This 'eastern population' could have come from anywhere between Helsinki and Yerevan, IMHO - an area 2000 k wide (!)

@ saman sistani said..."Dienekes I wish you were more active in the comments, your insight and knowledge is much appreciated."

Not to put words in Dienekes mouth, but i suspect he has said everything there is to say for the moment.

IMHO, there are still too many floating variables and 'blank spots' in aDNA data to prove or disprove anything.

As I mentioned in my first point, we need to test the Myceneans. There we have a 'fixed point'. A wide-covering and detilaed study using isotopes and genomic sequencing will produce pretty firm convictions.

@Rokus"Please don't confuse people -and yourself- even more. EEF is most of all WHG"

Please Rokus, The fact that you think Basal Eurasian is equivalent to Early Near Eastern farmer, let alone EEF shows me, you absolutely don't know what you are saying. "Basal Eurasian" is not equal to EEF and so is the rest of Stuttgarts ancestry not equal to WHG.

I have absolutely never come across a single knowledgable person who would claim that EEF is mostly WHG.

Effectively, not "completely" decoupled, from a population evolution viewpoint, and I included Italy rather than excluding it. The Carpathians and Tatra mountains and the Alps form a formidable barrier, and were largely glaciated during LGM. The Epigravettian of the Balkans is its own local development, and its range is well-established. There are very few settlement sites in eastern Central Europe (Austria, Czech Republic, Slovakia, Poland, Hungary) during LGM, and a lot of evidence for total abandonment during the late, dry phases. There are some temporary sites of Balkan origin in SW Romania (which is part of the Balkan peninsular, anyway), but no indication of permanent settlements during LGM, AFAIK.

Could there have been contact with people occupying habitable areas in the Ukraine? It is possible, but there is not much archaeological evidence for it, and moreover, the two climates were extremely different. In fact, the Balkans saw some of the mildest and most hospitable climate and most diverse fauna in all of Europe during LGM, and accordingly could host a fairly large population (as also evidenced by its many sites).

"There is no reason to assume that a complete merge between "Aurignacian+Gravettian WHG" related cultures during the Upper Paleolithic or Mesolthic."

Again, it doesn't have to be complete, but the Gravettian was a pan-European phenomenon, and the ancient DNA shows a pretty narrow WHG/ SHG cluster with just a moderate N-S cline - no evidence of two very different populations.

"I think it just reflects your antipathy for people from the Balkans"

Why would you make such a nonsensical statement? If I try to argue that the pre-neolithic people of the Balkans were closely related to NW Anatolians from which much of the European Neolithic derives, but not so much to WHGs, how does that reflect antipathy? I am elevating their status with respect to modern European origins.

Check out K=17 and beyond. Notice the teal component, which is present in half the Yamna is present in All Indo-European speaking populations, ancient samples and modern from Corded Ware, to Yamna, to modenr Northern Europeans, to modern Southern Europeans and Nearesterners, while the dark blue which makes up the other half of the Yamna component is not present at all in the Near East or Caucuses. One could argue the 'teal' is the uniting PIE component. We would expect the dark blue to be in the West Asian Indo-European speaking groups if the Yamna were the actual PIE.

I won't say anything for certain now, we need more genomes from the Near East and Caucuses IMO.

The rather low Yamnaya ancestry in the HungaryGamba_BA sample according to Figure 3 seems to call into question what I wrote about an expansion of R1b and Italo-Celtic from the Yamnaya in the Carpathian Basin. However that's not true upon closer inspection, because that sample has rather strong WHG ancestry which wasn't present in the preceding HungaryGamba_CA sample. The explanation is straightforward: The Yamnaya in the Carpathian basin differed from the Yamnaya of Samara by having much stronger WHG ancestry, which would be natural if it came from a more western part of the steppe.

There seems to be a connection between the above observations and the fact that BR1 has 0% of the Dodecad K12b Gedrosia component. Likewise, the surprisingly strong Yamnaya-like ancestry in Norwegians, Icelanders and Scots seems to be related with their strong Gedrosia component (European peak in Scots), also with their relatively strong MDLP World-22 Indo-Iranian component (European peak in Norwegians). So, if my above reasoning is correct, these components were not always associated in Europe with haplogroup R1b.

According to Figure 3, populations like the French and the Northern Spanish have more Yamnaya-like ancestry than the HungaryGamba_BA sample, and this seems to imply that a lot of this is actually from the central European Corded people, even though the latter were dominated by R1a. Apparently, this R1a got displaced by Bell Beaker R1b, but the autosomal Yamnaya-like ancestry of the Corded people was absorbed by the Bell Beaker people. Similar things happened in Scandinavia, where haplogroup I1 came to dominate, although autosomally there is still a lot of Corded Ware ancestry in Scandinavians.

Finally, I noted that Halberstadt_LBA, the sample from late Bronze Age Germany, again has somewhat stronger Yamnaya-like ancestry than the earlier Bell Beaker and Unetice. I suppose the most simple explanation for this is that the local Urnfield culture north of the Harz was related with the Lusatian culture to the east, which is also reflected in this individual's haplogroup, R1a-Z280.

"There are very few settlement sites in eastern Central Europe (Austria, Czech Republic, Slovakia, Poland, Hungary) during LGM, and a lot of evidence for total abandonment during the late, dry phases. There are some temporary sites of Balkan origin in SW Romania (which is part of the Balkan peninsular, anyway), but no indication of permanent settlements during LGM, AFAIK."

I just did a quick search on Google Scholar and found sites in Albania, Thessaly, Serbia, Epiros and Macedonia that indicate occupation in the Upper Paleolithic and Mesolithic. I've pasted some of the information below.

These sites would not have been completely cut off from Europe. For instance, Italy could easily have been reached from Thessaly in several ways:

1. Walking over the passes from the Balkan central plateau to the Adriatic, by way of Shishtavec, Lake Ohrid, Prespa Lakes, Samarina-Konitsa, and the Metsovo Pass, then up the shoreline of the Adriatic, which during the Mesolithic would have been a beach or grassy plain.

2. Walking up the central plateau, which extends from the Pelaponnese all the way to Hungary-Slovakia-the Czech Republic-Austria-Germany and the Ukraine. In fact, the exact path that Vlach shepherds still use. It really isn't that far (less than a thousand miles from Greece to Moldova).

So very improbable that the central Balkan plateau was isolated, especially given the abundance of water, game, and its ecozone continuity with the Alps.

An additional line of evidence that Greece and the Balkans were not cut off from the rest of Europe is the fact that wolves, the Eurasian brown bear, lynx and the ibex in the Dinaric Alps are not speciated from the Swiss and Austrian Alps. Even the Balkan chamois is considered to be a sub-species of the Chamois.

---------------------------

The Palaeolithic and Mesolithic of Albania: Survey and Excavation at the Site of Kryegjata B (Fier District)http://www.equinoxpub.com/journals/index.php/JMA/article/viewArticle/638

Late Mesolithic lifeways and deathways at Vlasac (Serbia)http://www.maneyonline.com/doi/abs/10.1179/0093469013Z.00000000070

The Neolithisation of the last hunter-gatherers in the Eastern Mediterranean between Western Anatolia and the head of the AdriaticMichael Templer, 2nd year PhD, Supervisor: Dr. Matthieu Honegger, Professor of Prehistoric Archaeology, University of Neuchâtel, Switzerland.

POST-GLACIAL TRANSFORMATIONS AMONG HUNTER-GATHERER SOCIETIES IN THE MEDITERRANEAN AND WESTERN ASIA -AMT MOORE - The Oxford Handbook of the Archaeology and …, 2014

The Palaeolithic of the Balkans. Proceedings of the XV World Congress of the InternationalUnion for Prehistoric and Protohistoric Sciences (Lisbon, 4–9 September 2006), Vol. 17,Session C33Andreas Darlas and Dušan Mihailović (eds.)Oxford: Archaeopress, 2008, 116 pp. (paperback), £29.00.ISBN-10: 1407303058.

I thank eurologist for having said that: "Effectively, not 'completely' decoupled, from a population evolution viewpoint, and I included Italy rather than excluding it". I based my theory of the Italian Refugium upon that. Probably we'll know the truth when these results will be published: "Genetic evidence ruled out one likely related group in the region, the Yamnaya, because their DNA showed the group had hunter-gatherer ancestry, which is inconsistent with the fact that two Indo-European groups, Armenians and Indians, don’t share it, Patterson said. That made Patterson look south, to the Maikop civilization, which likely had significant contact with the Yamnaya, as a plausible culture where Indo-European languages originated. Samples have been obtained from Maikop burial sites, but the DNA work to test that proposal is pending, Patterson said".

'Notice the teal component, which is present in half the Yamna is present in All Indo-European speaking populations, ancient samples and modern from Corded Ware, to Yamna, to modenr Northern Europeans, to modern Southern Europeans and Nearesterners, while the dark blue which makes up the other half of the Yamna component is not present at all in the Near East or Caucuses. One could argue the 'teal' is the uniting PIE component. We would expect the dark blue to be in the West Asian Indo-European speaking groups if the Yamna were the actual PIE.'First of all it should be determined where this 'Neolithic' component was really from. I think it is almost inconceivable that eastern European Neolithic influences would be so different from central Europe as to have much closer, or even quite different links to the Near East. Most of all, the continental Neolithic in east and west shared the same history. So maybe there rather is a link to the mysterious Gedrosian component instead? And how did this affect Samaria?Samaria was also the place where HG pottery (i.e. the Ceramic Mesolithic) first entered Europe. The same kind of pottery also emerges later in NE Europe with the mesolithic Swifterbant culture. All of this pottery is clearly linked with Funnel Beaker, Bell Beaker and Corded Ware pottery. Probably the teal/'East Neolithic'/Gedrodia component is one and the same, and goes back to Mesolithic times. The broader PIE homeland may have encompassed all of these enclaves that originally had an eastern origin. A pity the Swifterbant genome is still missing!

"Basal Eurasian" is not equal to EEFcorrect, like I already quoted from Lazaridis:'Stuttgart, fit as admixed with 47-79% Loschbour-related ancestry and 21-53% “basal Eurasian” ancestry', EEF is up to ~80% WHG.

'and so is the rest of Stuttgarts ancestry not equal to WHG.'Nobody will buy this 'logic'. Is it so difficult to admit you have it all wrong?

'I have absolutely never come across a single knowledgable person who would claim that EEF is mostly WHG.'Maybe wrong contacts, though anyway you should rather try to read the relevant scientific literature by yourself.

Sure, theoretically - but I don't think the timing works out. IMO other examples such as Uralic speakers and the combined Jewish diaspora suggest that a lot more than 2,000 - 3,000 years are required to extinguish an acquired cline from two significantly different populations. Moreover, a quickly multiplying population that expands into several directions will "freeze" the cline signature in combination, even if each subgroup starts to homogenize. In the case of the Early European Neolithic (EEN), the cline in PC2 is about 3 times as large as the one in PC1, across Europe, indicating that PC1 was homogenized at least several thousands of years before the samples (i.e., that long before 5,700 - 5,000 BCE, across Europe), while the N/S gradient of PC2 largely persisted.

Now, we have the early Neolithic Greece at 6900–6500 BCE, contemporaneous with NW Anatolia but with no cultural link other than the grain and pulse seeds and the animals, the Starčevo culture starting at around 6,000 BCE, and LBK starting to expand between 6,000 and 5,500 BCE. Similarly, Cardium Pottery starts along the Adriatic coast ~6,400 - 6,000 BCE, moving into S Italy ~6,000 BCE, and into Iberia ~5,500 BCE. So, something like a millennium or at best two overall, but virtually no time within the Balkans.

In other words, the population within the Balkans (and likely NW Anatolia) already had the characteristics of EEN (central, narrow PC1 and wide N/S cline in PC2) at and before the beginning of the Neolithic. They shared WHG from before LGM with the remainder of (W) Europe, but also some W Asian (Basal Eurasian?) - likely much more moderately so than current S Levantine populations.

Re: What I wrote yesterday: Maybe BR1 and BR2 are not indicative of the Yamnaya in the Carpathian Basin. BR1 lived several centuries after the Yamnaya incursion, and after the appearance of R1b in German Bell Beakers, moreover in the fully fledged Bronze Age, no longer in the Copper Age. I think it's possible that BR1 was partly descended from eastern Balkan people who may have had admixture from western steppe people predating Yamnaya. If that's the case, the German Bell Beakers may have had stronger and more regular Yamnaya ancestry, quite independently from Corded Ware admixture.

Yet, the Yamnaya-like ancestry in German Bell Beakers is too weak to explain Yamnaya-like ancestry in western Europe. They had less than the present-day Scottish! So either some heavily Corded Ware admixed Bell Beaker groups migrated West, or Iron Age steppe groups like IR1 had more influence than I expected.

Someone says: "That is actually not true, unfortunately. A few weeks ago, Nick Patterson stated it was a misunderstanding in the comments section of one of the posts at Eurogenes". I could say that this could be a good news for me...

Marnie writes: "These sites would not have been completely cut off from Europe. For instance, Italy could easily have been reached from Thessaly in several ways [...]".Of course I have always said that I am pretty sure that, beyond other Y and mt hgs, R1b1-L389+ (of course with other upstream and downstream subclades till perhaps R-L51) migrated from Italy to Iberia by sea 7500 years ago as Zilhao demonstrated, but whether they were from the Mesolithic Italy or come from the Balkans had to be demonstrated and it was an open road yet... Of course only aDNA will be able to give the last word. I bet upon Italy, but my acquired cousin from Tuscany, Fabrizio Federighi, R-M269, with the mutation PF7566 and PF7569 finds the unique link so far in a Greek, and that Cardial came from the Balkans and perhaps before from Asia Minor (someone would like from Lebanon before) is still an open question.

Your whole scenario, claim is so wrong I don't even know where to start. Before I can start a serious discussion you need to have the basic knowledge. Which I can't see at you.

Since when can a sample which is 20-50% Basal Eurasian, be considered as 80% WHG if the "Basal Eurasian" part is an independent component from WHG. I don't get your logic. It is so confusing that I don't even know what to say.

The problem with your whole "Logic" Rokus is that you think "WHG related" means WHG.

The whole point you missed is that Basal Eurasian is an independent component from WHG. And the Near Eastern farmer component is made up of Basal Eurasian and an WHG RELATED component ALREADY in the Near East. This is the Proto_Farmer component. Now the only question remaining is how much of WHG did these proto farmers catch up in the Balkans to become EEF. And even the Lazaridis paper concluded that it must be between (I don't know the exact number anymore) ~40 to ~2% Depending on how admixed we consider the Bedoin samples taken as refference. And Lazaridis in his own paper comes to the conclusion that both extreme ends are very unlikely and something in between the most logical.

If you follow the link above you'll see an ADMIXTURE analysis that is inclusive of both ancient and modern genomes all around the world.

Anyway, the ancient Yamna genomes are feature in the study. If you check out K=16 through K=19 you'll notice the Yamna basically come out 50% dark blue and 50% teal. The dark blue seems to peak in Paleolithic European hunter-gatherers while the teal peaks in the Caucuses.

In K=16 through K=19 I notice that although the dark blue is in Europe, even in Southern Europe, it lacks in the Caucuses and the Near East. While the teal is present in all Indo-European groups, from the Caucuses/Near East to all of Europe from North to South.

Basically, the teal component which peaks in the Caucuses seems to be the common uniting component in present Indo-Europeans, and there seems to be no evidence, based on this limited ADMIXTURE run, of Northern influence into the Near East or Caucuses. The Yamna were half Palelolithic-Hunter Gatherer yet there's no evidence of this component in the Near East or Caucuses from this ADMIXTURE run.

"I just did a quick search on Google Scholar and found sites in Albania, Thessaly, Serbia, Epiros and Macedonia that indicate occupation in the Upper Paleolithic and Mesolithic. I've pasted some of the information below."

...which are all Balkan sites and thus support what I have been saying.

"These sites would not have been completely cut off from Europe. For instance, Italy could easily have been reached from Thessaly in several ways:..."

I have always included the habitable areas of Italy - nothing new, here.

"So very improbable that the central Balkan plateau was isolated,..."

I agree - but that's a straw man. No one in their right mind would hold such a position.

Let's recapitulate: There was R1b1 in early Neolithic Spain and there was R1b1 in EHG in Samara. This means R1b1 was spread over a vast geographic area. We have no data from West Asia, but it's conceiveable that R1b1 was present there as well. Then, we have, a little closer to modern west European R1b, R1b1a in Yamnaya, and quite a bit closer, R1b1a2a. However, since there is obvious cultural and autosomal West Asian admixture in Yamnaya, we cannot rule out that these latter haplogroups came together with this admixture from West Asia. And if that really was the case, then we cannot rule out that R1b1a2a in German Bell Beakers merely shared the common West Asian source with Yamnaya, instead of being descended from it.

As has often been pointed out, the Bell Beaker people were often of a brachycephalic, planoccipital type with big, convex nose, reminiscent of modern-day West Asians. So it would seem plausible at first sight that their R1b1a2a had rather come from West Asia than from Yamnaya.

But it's revealing to study their results in the ADMIXTURE analysis. There it's undeniable that the Bell Beaker samples have less of the greyish West Asian component than the Corded Ware samples have, which in turn have less than Yamnaya has. So they were definitely not particularly West Asian admixed people. But they not only have absolutely less of it, but also relatively, in relation to the blue HG component. In Yamnaya the proportions between greyish West Asian and blue HG component are roughly 50 : 50. In Corded people the West Asian component is a little weaker than the blue HG component. And in Bell Beaker people the ratio HG : West Asian component is about 3:1. Now if we also look at BR1, the first of the HungaryGamba_BA samples, the same trend is even more accentuated. Here, the West Asian component is tiny compared to the blue HG component.

And I think here lies the solution to the puzzle of the apparently much too low Yamnaya ancestry in BA Hungary. The problem is that there is only one Yamnaya-like component in Figure 3, this distorts things. Conceivably, westernmost Yamnaya wasn't necessarily like Yamnaya in Samara. At the western end of the steppe, the EHG must have been closer to WHG, and there was less of the West Asian component. These groups migrated into the Carpathian Basin and mixed with the EEF locals, which resulted in people like BR1, with strong WHG component and modest „Yamnaya component“, also weak West Asian component in ADMIXTURE. And if you mix someone like BR1 with Corded people, you get something like German Bell Beaker people! This explains their stronger „Yamnaya component“ in Figure 3. That they did admix is quite a natural assumption. The PCA shows that they varied quite a bit in their Corded Ware admixture. Some were very close to BA Hungary, others clearly on the way towards Corded people. But since the Corded people had much more R1a than R1b, the R1b in Bell Beaker people came from another source, and that was probably the BR1-like people from the Carpathian Basin, who probably had it from Yamnaya. I have to stress here that BR1's West Asian component is very weak. In Dodecad K7b she has 0.01% West Asian, and in Dodecad Globe13 it's 0.00%. In the unsupervised ADMIXTURE run in this paper she does have a little bit of the West Asian component. But the fact remains, that the change from CO1 to BR1 was mostly one of increasing HG ancestry.

That the main source for German Bell Beakers was neither Corded-like, nor Samara-like, nor West Asian can also be seen in the fact that, according to table S7.1, Bell Beakers share the largest portion of common drift with KO1, the HG-like early farmer from Hungary. He was probably close to EHG in the westernmost steppe.

It's also interesting to note that, according to figure S9.8, German Bell Beakers can be modelled as a mix of Spain_MN, Esperstedt_MN and 24.2% Karelia_HG. This shows that the West Asian impact was not very important in them.

I think at least the absence of West Asian admixture in the Karelian EHG refutes the possibility that R1a1 had a recent West Asian origin connected to IE expansions. I don't know if West Asia was already very basal and distinct at the time when R1a originated. Of course it's possible to have a haplogroup without having any detectable admixture from the source population of this haplogroup, if it came from just a single, distant exotic ancestor. But I suppose hunter-gatherers usually migrated in groups, not as lonesome wolves, and hence admixture events introducing a new haplogroup would affect whole groups and many ancestors. If we compare it with haplogroup N, in populations where N is strong we always see some Siberian ENA admixture, even if it's just slight, it's still detectable, and I would expect the UP and Mesolithic spread of R1a to have been associated with autosomal admixture from it's source population as well.

Good point about the relative paucity of the West Asian component in BR1 and Bell Beaker. If the West Asian is weaker in W than E Yamnaya that would suggest that the West Asian component really isn't associated with Tripolye-Cucuteni (not that anyone seems to be arguing that anymore). Could still be Caucasian or Central Asian (or both).

Karelia_HG has R1a1*(xM17/M198). I have not seen any age estimate for R1a1-M459/SRY10831.2, but Y-Full currently puts upstream R1a-M420 at ~13700 BP, and R1a1a-M17 at ~8500 BP. That leaves a rather large window for R1a1 to potentially spread around the beginning of the Holocene.

We can't assume we'd recognize South Caspian forager ancestry from before the spread of the Neolithic, and there are Paleolithic and Mesolithic connections between north and south.

"Anyway, the ancient Yamna genomes are feature in the study. If you check out K=16 through K=19 you'll notice the Yamna basically come out 50% dark blue and 50% teal. The dark blue seems to peak in Paleolithic European hunter-gatherers while the teal peaks in the Caucuses."

Rather suggestively, the teal component also looks about as high among the Kalash as it is in the Caucasus. Many thanks.

@ eurologist "@Gioiello Good to see you back". I thank you, but I have to try writing not so much, because when I write a lot they always end banning me. I thank Dienekes who never banned me, but I wouldn't put him in difficulty, now that we are arriving in the solution of all our questions, even though strange things happen: a paper on Maikop annonced and soon retired, the Iberian R1b samples tested for M269, M35 (!), M69 (!) but not for L389 and V88...But labs and funds aren't ours...

"I agree - but that's a straw man. No one in their right mind would hold such a position."

Right.

I went back in thread and read your original statement. Here it is:

"Ötzi was found in a region that can easily be reached from the NW Balkans along several rivers (even via the Danube). I have previously argued that the ~Ice Age Maximum (LGM) Balkans were geographically and demographically decoupled from the rest of Europe (likely with the exception of habitable zones in Italy - especially the-then extended Po valley); but, because the Bosporus was a land bridge and because of known seafaring skills, they were very much connected with NW Anatolia. So, ~45-25 kya, they were all the same (old Aurignacian+Gravettian WHG with a cline to the Near East), and then you have a 15 ky gap to the beginning of the Neolithic. Perhaps this explains why the early Neolithic samples are so incredibly narrow in PC1 - that takes more than a few generations, especially in a large, quickly growing, and geographically widely expanding population. But in PC2 they are rather far apart, meaning that (as today) they were sorted by latitude during LGM and only thrown into similar heaps during the westward Neolithic expansion."

Regarding the statement, there is not reason to think that the Gravettian and Aurignacian genetic legacy in Europe all lumped together.

There is no reason to say there is a "15 thousand year gap" in the archaeological record in the Southern Balkans.

Regarding Oetzi, his DNA was found in the Alps, and if someone were to find a Mesolithic person from the Dinaric Alps (or the related sub-alpine region), the DNA from this person would most like look like Oetzi, only with WHG and "farmer" DNA, probably with more of a shift toward "farmer" DNA.

Archaeologically speaking, the adoption of farming in the Southern Balkans increasing does not look like a process of population replacement, but a local phenomena, with continued contact both between the European Alpine regions, the Black Sea low lands, and Asia Minor.

Indo-Europeans are not very present in the Caucasus. The latter is rather a stronghold of non-IE groups, with some exceptions. And the Near East & Anatolia inhabited mainly three groups of IEs (prior to Hellenism): Armenians, Thracians, Phrygians etc who were from the Balkans (in the case of the Armenians that's at least Herodotus' account), Indo-Iranians who came from the east, and the Hittites and relatives whose origin is disputed. It has often been maintained that Armenians completely lack any admixture from northern West Eurasians. In present-day Anatolian Turks and in several Indo-Iranian groups that type of admixture is more clearly present. And in this ADMIXTURE run I can in fact see the faint presence of the blue component also in Armenians, at K=16 and K=17 for instance.

Arch Hades "Basically, the teal component which peaks in the Caucuses seems to be the common uniting component in present Indo-Europeans, and there seems to be no evidence, based on this limited ADMIXTURE run, of Northern influence into the Near East or Caucuses. The Yamna were half Palelolithic-Hunter Gatherer yet there's no evidence of this component in the Near East or Caucuses from this ADMIXTURE run"

Thanks . That's is what I thought you were getting at. Certainly makes sense to me- if we are to look at a pan-IE signal; that's it ; and dovetails with dienekes earlier finds of west Asian component.

Simon Id be interested in your retort, in light of ur comments on Corded ware and bronze age Hungarian sample

"the Iberian R1b samples tested for M269, M35 (!), M69 (!) but not for L389 and V88... But labs and funds aren't ours..."@ DienekesOf course V35 and V69. They were so clerical errors that you could correct them. Unfortunately once posted, there isn't the possibility to correct anything. Anyway my discourse doesn't change. I hope that those samples are put at disposal of everyone like the 1KGP samples and everyone could look for any SNP tested. If I remember well neither this sample from Spain nor the P312 Bell Beakers from Germany were tested so before, and of course to answer my questions has a great impact upon the theories made so far.

Of course it may still seem possible to speculate that Bell Beaker R1b-P297 was originally associated with the tiny bit of West Asian admixture in people like BR1. But there is really no evidence for the idea that R1b-P297 originated in West Asia. R1b-P297 (xM269, M73) seems to be extremely rare, and its two subbranches M269 and M73 are much more common outside of West Asia than within. Their distribution looks like they had split somewhere on the Eurasian steppe, one branch going west, the other east. And we know that R1b-P297 was present in Yamnaya. So regardless of its ultimate origin, it seems most reasonable to assume that it spread with steppe people.

Regarding the spread of the West Asian autosomal component in Europe, I think there were two quite different kinds of migration behind that. On the one hand it spread with Yamnaya and related eastern European groups, starting in the Copper Age. Mainly because of this its really ubiquitous now, even in far northwestern Europe. But there was also another movement behind its spread, and this can be seen in the West Eurasian PCA, because it affected some populations that much that they are now inbetween Europeans and West Asians. These populations are Italy (including Sicily, but not Sardinia), Malta, and southeastern Europe (Albania, Greece, Bulgaria, Cyprus). And this spread of the West Asian component was quite evidently associated with y-haplogroup J2, as this is rather common in all these populations. This expansion seems to have started later than the aforementioned one. We've seen in Hungary (Gamba et al. paper) that J2 + additional West Asian admixture appeared eventually inbetween 2000 BC and 1200 BC. Also, Dienekes' Rolloff experiments showed that the admixture with the West Asians happened in Greece around 1840 BC and in Bulgaria around 1670 BC, perfectly in line with Gamba's evidence. I don't know what ethnic groups stood behind this spread. At least I note that in Italy and southeastern Europe non-IE languages survived long enough until there were scripts to document them. And in any case this Bronze Age J2 spread is too late to account for the Indo-Europeanization of Europe in general. (Because the time would simply not suffice. And there is no common IE word for bronze.) Moreover, at the edges of its spread, where it faded out, it's not clear why it should have entailed a language replacement. Some slight admixture doesn't necessarily entail language change, this would presuppose at least a dominating new elite.

@ Capra

The FTDNA R1a project has similar estimates, based on „Big Y“-results: R1a1a at 8000 BP and R1a1 at 12000 BP.

"There is no reason to say there is a "15 thousand year gap" in the archaeological record in the Southern Balkans."

Mamie,

Not a gap in records, a gap in the genetic exchange period, as far as meaningful on a population evolution measure. The Balkans and N Italy (minus the Alpine region) and the adjacent Anatolia were effectively isolated from the remainder of Europe.

"Googling on google scholar will quickly tell you that Italy and the Balkans formed one on [sic] the refuges for the Gravettian and Epi-Gravettian."

Which has been my point, all along.

"Both Italy and the Southern Balkans show very clearly that they were linked with the Alpine regions of Italy and Austria."

Sure, they were, as I have pointed out numerous times - but not around LGM, as I demonstrated above.

"You can fritter away the next hundred years of pop gen research trying to prove otherwise, but you won't be able to do so."

Not sure what that means. I am retired - so (i) I neither have 100 years left, nor (ii) do I understand what my association to "pop gen" is supposed to be.

"By the way, the name is Marnie."

As in, "My name is Luka?" (Not sure if I got that pop-gen reference right. I am sorry, I have been distracted - I am sitting in the morning at the diner on the corner, when I'm feeling someone watching me. There's a woman on the outside, and I'm trying not to notice that she's hitching up her skirt...)

As noted the extensive SNP testing in this study has results that are explosive. The authors used the rapidly expanding universe of SNPs found by more and more walk the Y tests.While some earlier results of I haploid group had as yet no living tests with the same SNPs or STRs, The I L161.1 L1498 found for subjects 0412 and 0172, has over 200 individuals with STR testing in the ftydna I2a group.

0412 is reported about 5310-5206 BCE, 0172 3360-3086 BCE

Similar interesting results may await in other samples of this report.

I am not an expert of hg. I of course, thus I ask you: from the YFull treeI-L161.1Y3753 * S2639 * Y7657... 33 SNPs9900 ybp, TMRCA CI 95% 77005500 ybp"formed 11300 ybp, TMRCA 6600 ybp I-L161.1* I-L1498S2643 * L1498/Y40085500 ybp, TMRCA CI 95% 72005000 ybp"formed 6600 ybp, TMRCA 6100 ybp I-L1498* id:YF01716it seems that I-L1498 has formed 6600 ybp and the MRCA lived 6100 ybp, thus did you mean ybp and not BC? If the dates are BC, YFull tree is underestimated of about 1500 yeas. From these 200 tested descendants you could say if these Y-s from Russia expanded to Western Europe or if happened the other way around. From the YFull tree seems that they live above all in the Isles, thus don't you think thar they, like probably the R1a and R1b, came from the Western European Refugia and expanded till the Russian plain?

The YFull experts are very smart and I am sure they reserve the right to correct their estimates in the future. Basically any MRCA is an estimated mean with some degree of uncertainty. You can ask YFull what that uncertainty is by their methodology for a start. Based on STR data, Ken Nordtvedt once had an estimate of about 12000 YBP for L161.1 and about 6000 YBP for L1498. The German site L1498 of about 5300 years ago is well within these best todate estimates of MCRA. BCE is the same as BC, CE means current era or 2015 years ago. Yes the older site 7325 YBP only has L161.1. A 2000 year error on any of these estimates would not be a surprise to me. There are L161.1 outlier present day tests from places like Thrace and Iraq. As to when, which direction, and with what y-DNA different expansions were I am waiting for more data.

Note that the YFull I-L161.1 "formed" estimate. ybp formed 11300 ybp, TMRCA 6600 ybp. Ken Nordtvedt has used STRs to predict clade and whether any living descendants have been tested. In I2a there are many subclades that are separated in STR space so that the time to divergence for a given archaic sample could be estimated. Hoping that future researchers will keep pulling out STR values as well as SNP.

Being 'too close' more likely means strong selection or rapid population growth. Both of which we expect to see due to known population growth. Migration waves should produce the opposite because they won't penetrate everywhere and there will be many refugeums.

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